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Lepiota maculans, an Unusual Mushroom Rediscovered after 105 years

Authors:
Joshua Birkebak at North American Mycological Association
Joshua Birkebak
  • North American Mycological Association
Else Vellinga
Else Vellinga
Ana Esperanza Franco Molano at University of Antioquia
Ana Esperanza Franco Molano
Michael G. Wood
Michael G. Wood
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Abstract and Figures

During a survey of macrofungi in the Great Smoky Mountains National Park in the summer of 2009, a species of Lepiota was recorded. This specimen is distinct in having pink to salmon to orange colors on the fruiting body and a pink spore deposit. After comparison with type material and a survey of the literature, we conclude this species is L. maculans Peck, described originally and known only from one collection in Missouri. Phylogenetic analysis of DNA sequence data supports placement within Lepiota of the Agaricaceae despite possessing a pink spore deposit. A complete morphological description, a description of the holotype, illustrations, and photographs are presented. This is the first report of L. maculans since its original description in 1905.
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Lepiota maculans, an Unusual Mushroom Rediscovered after 105
years
Author(s): Joshua M. Birkebak, Else C. Vellinga, Ana E. Franco-Molano, Michael G.
Wood and P. Brandon Matheny
Source: Southeastern Naturalist, 10(2):267-274. 2011.
Published By: Humboldt Field Research Institute
DOI: 10.1656/058.010.0207
URL: http://www.bioone.org/doi/full/10.1656/058.010.0207
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SOUTHEASTERN NATURALIST
2011 10(2):267–274
Lepiota maculans, an Unusual Mushroom Rediscovered
after 105 years
Joshua M. Birkebak1,*, Else C. Vellinga2, Ana E. Franco-Molano3,
Michael G. Wood4, and P. Brandon Matheny1
Abstract - During a survey of macrofungi in the Great Smoky Mountains National Park
in the summer of 2009, a species of Lepiota was recorded. This specimen is distinct in
having pink to salmon to orange colors on the fruiting body and a pink spore deposit. Af-
ter comparison with type material and a survey of the literature, we conclude this species
is L. maculans Peck, described originally and known only from one collection in Mis-
souri. Phylogenetic analysis of DNA sequence data supports placement within Lepiota
of the Agaricaceae despite possessing a pink spore deposit. A complete morphological
description, a description of the holotype, illustrations, and photographs are presented.
This is the ¿ rst report of L. maculans since its original description in 1905.
Introduction
Lepiota (Agaricaceae, Agaricales) is a large genus (estimated to have at
least 500 species worldwide) of pale-spored, saprotrophic fungi that are poorly
known in North America (Vellinga 2004). Most monographic treatments of the
group (Kauffman 1924; Morgan 1906a, 1906b, 1906c, 1907; Murrill 1914) are
outdated as they lacked information on many pertinent (particularly microscop-
ic) details, and only one has focused on a portion of the southeastern United
States (Akers 1997).
During a macrofungal diversity survey of the Great Smoky Mountains
National Park by researchers affiliated with the University of Tennessee,
University of California at Berkeley, University of Washington, Mycological
Society of San Francisco, and British Mycological Society, a distinct Lepiota
specimen was found with a predominantly pink and orange coloration. Af-
ter further study, the identity was determined to be Lepiota maculans Peck,
originally published from a collection made in Missouri (Peck 1905). To our
knowledge, this fungus has not been reported since its original publication. A
cursory examination of 79 undetermined specimens of Lepiota at the Univer-
sity of Tennessee herbarium was conducted in search of other collections of
this species. Despite the extensive collection of fungi from the Great Smoky
Mountains National Park (Hesler 1937, Lickey et al. 2007, Petersen 1979), no
additional specimens of L. maculans were found.
1Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville,
TN 37996. 2Department of Plant and Microbial Biology, University of California, 111
Koshland Hall, Berkeley, CA 94720-3102. 3Laboratorio de Taxonomía de Hongos, Insti-
tuto de Biología, Universidad de Antioquia A.A.1226, Medellín, Colombia. 414856 Sat-
urn Drive, San Leandro, CA 94578-1349. *Corresponding author - jbirkeba@utk.edu.
Southeastern Naturalist Vol. 10, No. 2
268
Methods
Morphological examination
Color notations of fruitbodies were taken from Ridgway (1912). Microscopic
characters were examined from dried material revived in 5% KOH. Spores for
measurement were taken from the spore deposit, and 20 cells or spores were mea-
sured for each character. All measurement summaries are given in the following
format: minimum–mean–maximum value. A Q-value is the ratio of cell/spore
length to cell/spore width.
Molecular examination
DNA extraction. A dried tissue sample between 10–20 mg was excised and
ground in liquid nitrogen with a micropestle and a pinch of sand in a 1.5-ml
microtube. DNA extraction was performed initially with an E.Z.N.A.® Fungal
DNA Kit (Omega Bio-Tek, Norcross, GA). This procedure yielded no PCR
products so a high-performance kit, E.Z.N.A.® HP Fungal DNA Kit (Omega
Bio-Tek), was used to remove polysaccharides and other potential interfering
factors. The isolated genomic DNA was diluted in two successive 1:10 sterile-
water dilutions.
PCR. Primers ITS1F (Gardes and Bruns 1993) and ITS4 (White et al. 1990)
were used to amplify the ITS1-5.8S-ITS2 region (a commonly used barcode
region in molecular fungal systematics) on a Bio-Rad C1000 thermal cycler (Bio-
Rad, Hercules, CA). A mixture of sterile water and 5X buffer, GoTaq, and dNTPs
supplied by Invitrogen Corp (Carlsbad, CA) was prepared for each dilution of
DNA and controls following manufacturer protocols. Our PCR protocol followed
that of White et al. (1990). PCR products were visualized on a 1.0% agarose gel
prepared with ethidium-bromide and a UV transilluminator. The ampli¿ ed ITS
product of L. maculans was cleaned using a QIAquick PCR puri¿ cation kit (Qia-
gen, Valencia, CA).
Sequencing. A sequence reaction was performed on the puri¿ ed ITS product
using a BigDye Terminator 3.1 Cycle sequencing kit (Applied Biosystems, Foster
City, CA). The sequence reaction solution was puri¿ ed with a Sephadex G-50
column (General Electric Healthcare, Piscataway, NJ) using separator strips
manufactured by Princeton Separations (Freehold, NJ). Sequencing was per-
formed on an ABI 3730 48-capillary electrophoresis genetic analyzer (Applied
Biosystems) at the Molecular Biology Resource Facility at the University of Ten-
nessee. Sequence chromatograms were inspected and edited using Sequencher
4.9 software (Gene Codes Corp, Ann Arbor, MI).
Phylogenetic analysis. The nrITS sequence was compared with those already
present in GenBank using the program BLAST (Altschul et al. 1990), and based
on the outcome a database of sequences from Lepiota sect. Lepiota was compiled.
The nrITS sequences were aligned with the program MAFFT version 6 (Katoh
and Toh 2008, Katoh et al. 2002). The sequence data matrix was analyzed by
maximum likelihood (ML) using RAxML version 7.2.3 (Stamatakis et al. 2008).
100 rapid ML bootstraps were performed. Lepiota ochraceofulva P.D. Orton and
L. elaiophylla Vellinga & Huijser were chosen as outgroup species. These species
J.M. Birkebak, E.C. Vellinga, A.E. Franco-Molano, M.G. Wood, and P.B. Matheny2011 269
differ from the subject taxon by having non-fusiform spores and a different type
of pileus covering.
Results
Taxonomy
Lepiota maculans Peck, Bulletin of the Torrey Botanical Club 32(2):77
(1905) (Fig. 1).
Pileus. 3.2–4.0 cm broad, plano-convex often with a slight umbo; margin de-
curved at ¿ rst, becoming straight with expansion, somewhat sulcate striate, often
eroded with age; disc “Dresden Brown”, “Ochraceous-Tawny” to “Buckthorn
Brown”, velutinous, breaking up in scales near center of pileus, scales sparse,
exposing context; context “Mustard Yellow” to “Amber Yellow” near the center,
“Salmon-Buff” to “Light Salmon-Buff” to “Orange-Pink” outward, at very edge
discolored “Peach Red” to “Scarlet”.
Lamellae. Free, nearly crowded, thin, 4–6 mm broad, “Pale Ochraceous-Buff”
when young, becoming “Salmon Color” to “Orange-Pink”, often discoloring
“Peach Red” to “Scarlet” near margin of pileus.
Stipe. 36–54 mm long, 2.5–3.0 mm wide at apex, gradually enlarging down-
ward or sometimes bulbous to clavate, 3–5 mm wide at the base, “Maize Yellow”
to “Buff-Yellow” at apex, “Mustard Yellow” on the lower half with numerous
À occose-¿ brillose patches; partial veil leaving a loose median annular zone.
Pileus covering. A trichoderm of erect, cylindrical, slightly yellow-brown pig-
mented, 117-188-345 × 7-9-11 m elements with obtuse to slightly acute apices,
commonly secondarily septate, arising from numerous short, cylindrical to clav-
ate elements near the base, though some elements have an intermediate length
(Fig.2).
Figure 1. Fresh basidiocarps of Lepiota maculans (collection JMB 05-08-09-18). Scale
bar = 1 cm. Photograph © M.G. Wood.
Southeastern Naturalist Vol. 10, No. 2
270
Spores. 8.8-9.5-11.3 × 4.4-4.9-5.4 m, Q-value 1.86-1.93-2.11, smooth,
oblong amygdaliform in pro¿ le, fusiform in frontal view, dextrinoid, not metach-
romatic in cresyl blue (Fig. 2).
Basidia. 22-27-32 × 8-9-11 m, clavate to subcylindrical, with four sterigmata.
Cheilocystidia. 13-19-29 × 6.0-7.5-11 m, clavate to subcylindrical, some-
times clavate pedicellate, occasionally arising sympodially (Fig. 2).
Clamp connections. Present in all tissues.
Habitat. Gregarious, terrestrial, in unkempt grass near edge of deciduous
forest.
Specimens examined. TENNESSEE: Cosby, GSMNP, 35°45'39"N,
083°12'38"W, elevation 635 m, leg. J.M. Birkebak, 5-Aug-2009, JMB 05-08-09-
18, TENN064381.
Figure 2. Micromorphological characters: PP = pileus covering, CC = cheilocystidia, S =
spores. (from JMB 05-08-09-18).
J.M. Birkebak, E.C. Vellinga, A.E. Franco-Molano, M.G. Wood, and P.B. Matheny2011 271
Comments
This species is set apart in the genus Lepiota by its distinctive coloration.
Many species of the related genus Leucoagaricus discolor orange or red, but no
other known members of the genus Lepiota discolor so strongly orange and pink.
It is unclear whether this coloration change is a staining reaction or a gradual
discoloration with age. Peck remarked in his commentary of the protologue that
L. maculans is easily recognizable due to the bruised À esh that turns reddish and
the lamellae that become reddish or pink upon drying. Ideally, future observa-
tions of fresh material should resolve whether the nature of discolorations is
speci¿ cally due to bruising or by maturation of the sporocarps.
Lepiota maculans is the only known species of Lepiota to have a "Seashell
Pink" spore deposit as opposed to a white or cream deposit for the remainder
of the species in the genus. The remaining characters, viz., pileipellis struc-
ture, spore morphology, and the presence of clamp connections, are typical for
the genus.
Molecular phylogenetic reconstruction using ITS sequence data suggests a
placement of L. maculans in Lepiota sect. Lepiota near L. ignivolvata Bousset &
Joss. described originally from France (Fig. 3). Both species have oblong spores
and a pileus covering composed of both long and short elements. Lepiota igniv-
olvata, however, is a robust species with a stipe that discolors orange in its basal
part when touched. This species is known only from Europe, where it has been
reported widely in numerous À oras (Horak 2005).
A study of the holotype (Glatfelter s.n.) at NYS re-af¿ rms our identi¿ cation
of L. maculans. Importantly, the spores of the type are consistent in shape, size,
and dextrinoid and non-metachromatic walls as the Tennessee material. Unfor-
tunately, due to the poor preservation of the type overall, we could not observe
cheilocystidia and trichodermial elements of the pileus covering, although an
underlying hymeniderm to the pileipellis was observed (as in our material). As
complete a description as possible of the holotype is presented below. The fol-
lowing microscopic observations are our own.
Pileus. 1.5–2 cm diam, thin, convex, subumbonate, dry, minutely and densely
squamulose, reddish-yellow, the center darker.
Lamellae. Broad, subdistant, free, white, gradually changing to red or pink.
Stipe. About 5 cm long, 2–3 mm thick, equal, tough, À occose or ¿ brillose,
hollow, whitish or yellowish, the annulus slight, evanescent.
Spores. 7.5–9.5 x 3.7–4.7 m, ovoid to broadly ellipsoid in pro¿ le view,
weakly dextrinoid, not metachromatic in cresyl blue.
Basidia. 27–35 x 9–10 m, clavate, thin-walled, 4-strigmate.
Pleurocystidia. Absent.
Cheilocystidia. Not observed.
Hymenophoral trama. Regular composed by cylindrical to inÀ ated hyphae up
to 18 m broad.
Suhhymenium. Pseudoparenchymatous, up to 20 m thick.
Southeastern Naturalist Vol. 10, No. 2
272
Pileal trama. Loosely interwoven, composed of cylindrical, radially arranged,
thin-walled, 3–12 m broad hyphae with hyaline or with yellowish red content in
KOH.
Pileus covering. Appearing as a hymeniform layer composed of clavate termi-
nal cells, yellowish in KOH.
Stipitipellis. Of pararllel and vertically oriented elements up to 15 m broad,
with terminal cells similar to those of the pileipellis and forming tufts more abun-
dant at the apex.
Clamps connections. Present in all tissues.
Figure 3. Maximum Likelihood phylogram recovered for the nrITS region using
RAxML version 7.2.3 for selected taxa in Lepiota sect. Lepiota from North America
and Europe, including one other species from the GSMNP, with bootstrap values indi-
cated at branch nodes.
J.M. Birkebak, E.C. Vellinga, A.E. Franco-Molano, M.G. Wood, and P.B. Matheny2011 273
Discussion
Basic knowledge of mushroom-forming fungi is relatively poor compared to
that of plants and animals. The fruitbodies of mushroom-forming fungi are often
ephemeral, and the taxonomy of many species relies on morphological traits
necessary to observe in fresh condition. Basic information such as distribution,
ecology, and phenology is at best preliminary, if not incomplete, for many fungi
of North America, including the southeast United States (Bessette et al. 2007,
Weber and Smith 1985).
Extensive survey work documenting fungal biodiversity in the Great Smoky
Mountains National Park for the All Taxa Biotic Inventory has been conducted
(e.g., Lickey et al. 2007). As of 2009, 833 new records for the Park have been
made, 75 of which are species new to science, adding to the 2550 previously
reported species to increase the total of known fungi in the Park to 3383 (http://
www.dlia.org/atbi/new_science/discoveries.shtml). Lepiota maculans represents
yet another new record to the Park. It is possible that L. maculans has been pre-
viously overlooked, but this seems unlikely given its striking coloration and the
intensive collecting efforts in the Great Smoky Mountains National Park. It is
more likely the species is rare or infrequent given its currently known distribu-
tion—Missouri and eastern Tennessee.
Acknowledgments
We are grateful to the Great Smoky Mountains National Park and Dr. Karen Hughes
at the University of Tennessee for providing collecting permits. We would like to thank
David Pratt and staff af¿ liated with the University of Tennessee Biology Field Station for
providing ¿ eld facilities, as well as Dr. Steve Trudell and an anonymous reviewer for pro-
viding helpful reviews on this manuscript. Partial funding by NSF grant DEB-0618293 to
E.C. Vellinga and DEB-0949517 to P.B. Metheny is gratefully acknowledged.
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The genus Auricularia comprises 10–15 recognized species worldwide, and most are considered to have intercontinental to cosmopolitan distributions. Though regional field guides for the southeastern United States treat only one or two species, five species, A. fuscosuccinea, A. auricula-judae, A. mesenterica, A. polytricha, and A. delicata , have been recorded from the region. This study seeks to evaluate and revise current species concepts in Auricularia using phylogenetic and morphological methods to better understand the species occurring in the southeastern United States. Historical collections from herbaria and fresh material from the field have been examined and sequenced at two loci, ITS and rpb2. Phylogenetic results indicate several diverse clades are in need of taxonomic revision. Previous reports of A. auricula-judae in the southeastern U.S. likely represent a clade of A. americana . Collections identified as A. delicata are found to be polyphyletic and distributed in four clades. Variation in the presence or absence of a medulla layer in some species, previously a key morphological character, has made it evident that additional characters are required to reflect the genetic diversity in the genus. A new taxonomic character, the schizomedulla, is discussed and shown to distinguish two novel species, A. subglabra and A. scissa , from the morphologically similar species A. delicata . A reticulate, merulioid hymenial surface can no longer be considered a character unique to A. delicata . Furthermore, ITS data from the voucher-specimen from which the genome of A. delicata was produced indicates this species is A. subglabra . The taxonomy and nomenclature of Peziza nigricans is discussed and it is shown to be the earliest priorable name for A. polytricha sensu auct. amer. , and the new combination Auricularia nigricans is proposed. We now recognize the following species of Auricularia from the southeastern U.S.: A. americana, A. fuscosuccinea, A. mesenterica, A. nigricans, and A. scissa.
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... Its spores generally resemble those of its sister species, L. subgracilis, and the type species of Lepiota, L. clypeolaria (Fig. 1). Lepiota viridigleba also bruises orange or pale red when handled and this feature is similar to the related species Lepiota ignivolvata and L. maculans (Fig. 1) (Birkebak et al. 2011). Unlike most species within section Lepiota, where clamp connections are commonly observed, L. viridigleba apparently lacks clamp connections in all tissues. ...
Phylogenetic analysis of rDNA sequences indicates that the sequestrate Amogaster viridiglebus is derived from within the agaricoid genus Lepiota (Agaricaceae)
Article
Full-text available
  • Feb 2013
The rare sequestrate fungus Amogaster viridiglebus is known from only one collection in California where it was discovered among Populus roots. Based on sporocarp coloration and spore morphology, this sequestrate taxon was putatively considered to be an ectomycorrhizal member of the Boletales. However, no molecular data were previously available to definitively determine the closest relatives of this fungus. Here we revisit the morphology of Amogaster viridiglebus and present a phylogenetic analysis based on ITS and 28S ribosomal DNA. Our phylogeny indicates that Amogaster viridiglebus is nested in the genus Lepiota, suggesting that this rare species has a saprobic trophic mode and does not form ectomycorrhizae with plants. A new combination, L. viridigleba, is made based on these phylogenetic results.
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Lepiota baiyunensis sp. nov. (Agaricales, Agaricaceae) from Baiyun Mountain, China
Article
  • Jul 2023
A novel species of Lepiota sect. Ovisporae from Baiyun Mountain, Guangzhou, China is described, based on phylogenetic analysis of both internal transcribed spacer region (ITS) and multi-loci (ITS, LSU, IGS and mtSSU) combined with morphological examinations. Lepiota baiyunensis sp. nov. differs from Lepiota lilacea by its light yellow lamellae, stipe covered with lamellar squamules, lacking true annulus, usually clavate cheilocystidia and trichodermal pileus covering composed of long, cylindrical elements.
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Lepiota sanguineofracta (Basidiomycota, Agaricales), a new species with a hymeniform pileus covering from Italy
Article
Full-text available
  • Aug 2013
Lepiota sanguineofracta sp. nov., a taxon with a hymeniform pileus covering, from Italy, is here described. A full description, colour pictures of basidiomata, line drawings of microscopic features, and ITS phylogenetic analysis are provided. It is morphologically and molecularly close (sister) to Lepiota coloratipes from which it differs mainly by olivaceus tinges on the pileus surface, basidiome surfaces and context strongly reddening on handling, a sweetish smell of withered rose and binucleate, not metachromatic spores.
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The Mushroom TWiG: A Marvelous Mycological Menagerie in the Mountains
Article
Full-text available
  • Dec 2007
We present an update on efforts to catalog the basidiomycete taxa, particularly the mushroom-forming fungi, of Great Smoky Mountains National Park (GSMNP) for the All Taxa Biodiversity Inventory. The goals of this project are to: 1) collect, identify, and voucher specimens with the help of visiting mycologists and volunteers; 2) extract DNA, amplify and sequence the nrITS region for barcoding, and deposit these sequences on GenBank; and 3) create species web pages for general public use. At present (April 2006), approximately 2000 specimens comprising about 770 species have been collected. As many as 45% are new Park records, and several may represent species new to science. DNA has been extracted from about 1000 specimens, and the nuclear ribosomal ITS region has been amplified and sequenced for about 500 of those. A surprising amount of genetic heterogeneity has been found, in part due to population migration patterns in response to glacial cycles. Studies with Artomyces pyxidatus support this hypothesis, showing distinct contributions from Central America and a second unidentified refugium.
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Ecology and Distribution of Lepiotaceous Fungi (Agaricaceae) A Review
Article
Full-text available
  • May 2004
Lepiotaceous fungi form relatively fragile basidiocarps with white, rarely coloured, spores. Most are saprotrophic forest-floor dwellers that grow in the lower litter layers of the soil, and probably decompose lignin and cellulose. They occur worldwide, with many representatives in tropical and temperate regions, and a few species in arctic-alpine areas and in deserts. Most taxa are agaricoid, though a relatively small number of secotioid variants exist. Because of their relative rarity, the clustering together of many species in limited habitats, and the differences in species composition along latitudinal transects, lepiotaceous fungi may be vulnerable to changes in the environment, both on a local (habitat destruction) and on a global scale (climate change). Sister taxa occur in different parts of the temperate zones of the Northern Hemisphere, indicating that vicariance events might have played a role in speciation. A few species have a very extended distribution, and those species occur either in man-made habitats or else in cooler habitats. Many ecological features of the lepiotaceous fungi are unknown, including the survival rates and colonization success of spores, nutrient and temperature requirements, longevity and size of genets. Conservation of existing diversity calls for policies underpinned by new ecological research, more taxonomical studies, and more recording projects.
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Katoh K, Misawa K, Kuma KI, Miyata T.. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Res 30: 3059-3066
Article
  • Jul 2002
  • NUCLEIC ACIDS RES
A multiple sequence alignment program, MAFFT, has been developed. The CPU time is drastically reduced as compared with existing methods. MAFFT includes two novel techniques. (i) Homo logous regions are rapidly identified by the fast Fourier transform (FFT), in which an amino acid sequence is converted to a sequence composed of volume and polarity values of each amino acid residue. (ii) We propose a simplified scoring system that performs well for reducing CPU time and increasing the accuracy of alignments even for sequences having large insertions or extensions as well as distantly related sequences of similar length. Two different heuristics, the progressive method (FFT‐NS‐2) and the iterative refinement method (FFT‐NS‐i), are implemented in MAFFT. The performances of FFT‐NS‐2 and FFT‐NS‐i were compared with other methods by computer simulations and benchmark tests; the CPU time of FFT‐NS‐2 is drastically reduced as compared with CLUSTALW with comparable accuracy. FFT‐NS‐i is over 100 times faster than T‐COFFEE, when the number of input sequences exceeds 60, without sacrificing the accuracy.
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New Species of Fungi
Article
  • Jan 1879
  • Bot Gaz
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MAFFT: A novel method for rapid multiple sequence alignment based on fast Fourier transform
Article
  • Jul 2002
A multiple sequence alignment program, MAFFT, has been developed. The CPU time is drastically reduced as compared with existing methods. MAFFT includes two novel techniques. (i) Homo logous regions are rapidly identified by the fast Fourier transform (FFT), in which an amino acid sequence is converted to a sequence composed of volume and polarity values of each amino acid residue. (ii) We propose a simplified scoring system that performs well for reducing CPU time and increasing the accuracy of alignments even for sequences having large insertions or extensions as well as distantly related sequences of similar length. Two different heuristics, the progressive method (FFT-NS-2) and the iterative refinement method (FFT-NS-i), are implemented in MAFFT. The performances of FFT-NS-2 and FFT-NS-i were compared with other methods by computer simulations and benchmark tests; the CPU time of FFT-NS-2 is drastically reduced as compared with CLUSTALW with comparable accuracy. FFT-NS-i is over 100 times faster than T-COFFEE, when the number of input sequences exceeds 60, without sacrificing the accuracy.
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Checklist of Odonata from Great Smoky Mountains National Park
Article
  • Dec 2007
The fauna and flora of Great Smoky Mountains National Park is being systematically studied and documented for the first time as part of the Smokies' All Taxa Biodiversity Inventory (ATBI). With direction from scientific authorities and Park staff, a team of citizen volunteers has undertaken a survey of odonates (dragonflies and damselflies). The survey is focused on adults and includes curated specimens, catch-and-release records, and reliable sight identifications. To date, 93 taxa (63 dragonflies, 30 damselflies) are reported from the Park. However, the habitat-, geographic-, and temporal-survey coverage is far from complete, and records from neighboring areas suggest the Park may contain more than 130 odonate species. All of the information is being stored in the online ATBI database.
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